Age and growth estimates of the bull shark,Carcharhinus leucas,from the northern Gulf of Mexico

Synopsis Length at age and growth rates for 59 bull sharks, Carcharhinus leucas, collected from the northern Gulf of Mexico were estimated from the band patterns formed seasonally in the vertebral centra. The combined age at length data for both sexes were applied to a von Bertalanffy growth model producing parameter estimates of L = 285 cm TL, K = .076, t₀ = –3.0 yr. Lengths at age for males and females were similar except that males did not attain as great a length as females. Growth was apparently slow and varied among individuals, but in general, was estimated to be 15–20 cm yr^–1 for the first five years, 10 cm yr^–1 for years 6–10, 5–7 cm yr^–1 for years 11–16, and less than 4–5 cm yr^–1 thereafter. Males mature at 210–220 cm TL or 14–15 yr of age; females mature at>225 cm TL or 18+ yr of age. The largest male (245 cm TL) was 21.3 yr old; the largest female (268 cm TL) was 24.2 yr old.     

Age and growth of the bull shark (Carcharhinus leucas) around Reunion Island,South West Indian Ocean

Sharks exhibit varied demographic strategies depending on both the species and the population location, which make them more or less vulnerable to fishing. Accurate evaluation of local age and growth parameters is therefore fundamental for the sustainable management of their stocks. Although demographic parameters have been assessed for bull shark (Carcharhinus leucas) populations in several locations of the world, this information is missing so far around Reunion Island, in the south-west Indian Ocean. To fill this gap of knowledge, age and growth data were gathered from the vertebrae of 140 individuals of C. leucas (77 females and 63 males, mostly adults) fished around the island between 2012 and 2019. After verification of the annual deposition of growth band pairs on these structures using relative marginal increment analysis on 40 individuals, band pairs were counted along the vertebral centrum for each individual. Thanks to this approach, growth was shown to significantly differ between male and female C. leucas around Reunion Island, with respective von Bertalanffy growth model equations of L_t =  and L_t = . Indeed, the females of the species fished in this area were significantly (P < 0.001) larger than local males, with an estimated difference in size of ~16.1 cm at 20 years old. They also apparently reach older ages, with an estimated maximum age of 33.50 years, against 29.75 years only for the males. The estimated size at birth around the island is larger than elsewhere in the world, varying from 92.30 to 100.00 cm depending on the method used. These results confirm that the population of C. leucas around Reunion Island exhibits a K-selected strategy, which makes it highly vulnerable to fishing pressure.     

Bioenergetics of the bull shark,Carcharhinus leucas,maintained in captivity

Understanding the energy requirements for captive sharks is important for their successful long-term maintenance. This information is critical in assessing the health of the animals and the suitability of their environment. We studied five bull sharks (Carcharhinus leucas) for up to 7 years in a 2.5 × 10⁶ liter oceanarium. Individual animal feedings provided information for food intake analysis. During the first 3 years, fork length increase was estimated to have averaged 1.9 cm/month (s.d. = 0.1), or 23.0 cm/year. Biannual measurements, begun in the fourth year, showed that growth rates decreased during the next 4 years to a mean rate of 0.6 cm/month (s.d. = 0.2), or about 7.0 cm/year. Mean food consumption from June 1988 to December 1992 was 3.4% body weight per week. Caloric conversion of weights incorporated into a simple bioenergetics model providing mean metabolic expenditures per animal was 5.7 (s.d. = 0.3) and 4 (s.d. = 0.5) kcal/kg/day for 1991 and 1992, respectively. © 1994 Wiley-Liss, Inc.     

Reproductive biology of the crocodile shark Pseudocarcharias kamoharai

From February 2005 to September 2007, a total of 490 crocodile sharks Pseudocarcharias kamoharai, caught as by‐catch in the swordfish and tuna longline fishery that operates in the tropical western Atlantic Ocean, was studied in regard to their reproductive biology. Maximum observed total lengths (L_T) were 1220 and 1090 mm for females and males respectively, with a high proportion of the catch being composed of mature specimens. Sexual maturity was attained at 760–810 mm L_T for males (L_T50 = 800 mm) and 870–980 mm L_T for females (L_T50 = 916 mm). The size at birth was estimated at 415 mm L_T. Temporal variation in gonad morphology and mass suggests that in this region P. kamoharai, an aplacental viviparous species with oophagy, does not show a well‐defined reproductive seasonality, with mating and parturition occurring possibly over an extended period of the year. Mean ±s.d . fecundity was estimated to be 3·9 (± 0·6) pups per reproductive cycle.     

Reproductive cycle of the blacknose shark Carcharhinus acronotus in the Gulf of Mexico

The reproductive periodicity of blacknose sharks Carcharhinus acronotus in the northern Gulf of Mexico was determined by examining reproductive tissues from specimens collected monthly from 2002 to 2005. Monthly changes in shell gland mass, right ovary mass and ovarian follicle diameter were assessed for 74 mature females. Temporal changes in testes mass, testes width and proportion of mature spermatocysts were examined for 64 mature males. Trends in female reproductive tissues suggested an annual peak in reproductive activity during June and July, while trends in male variables suggested an annual reproductive peak during May and June. Although male and female reproductive activity peaked in different months, a strong synchronicity existed between the proportion of mature spermatocysts and the diameter of the largest ovarian follicle. Based on these results, the mating season of blacknose sharks lasts from mid-May to July in the Gulf of Mexico. Maximum embryo sizes were observed in May, which suggested that partition occurs during late May or early June. Results indicate that blacknose sharks have a clearly defined annual cycle in the Gulf of Mexico. This conclusion is further supported by the complete absence of gravid females without vitellogenic ovarian follicles among all mature females examined.     

Assessing multiple paternity in three commercially exploited shark species: Mustelus mustelus,Carcharhinus obscurus and Sphyrna lewini

In this study, multiple paternity (MP) was investigated in three commercially important shark species, common smoothhound Mustelus mustelus, dusky shark Carcharhinus obscurus and scalloped hammerhead Sphyrna lewini occurring in southern Africa. Reduced marker panels of between five and six microsatellite loci were constructed for each species and used to genotype and assess the presence of MP in a total of 60 M. mustelus individuals from six litters, 90 C. obscurus individuals from 14 litters and 54 S. lewini individuals from 13 litters. Analysis in GERUD and COLONY revealed the presence of MP in all three species. Multiple paternities were observed in 67, 35 and 46% of the litters of M. mustelus, C. obscurus and S. lewini, with corresponding average sire size of 1·6, 1·4 and 2·0, respectively. The variation in the rate of MP among the three species is in accordance with previous studies whilst the comparatively high frequency of MP observed for M. mustelus, matches what has previously been reported for shark species demonstrating aggregation behaviour.     

First evidence of multiple paternity in the blue shark (Prionace glauca)

Multiple paternity (MP) is defined as the behaviour in which females successfully mate with multiple males leading to offspring from different sires within the same litter. MP seems to be frequent and an evolutionary advantage in elasmobranchs. Here the authors report for the first time the occurrence of MP in the cosmopolitan blue shark Prionace glauca L. The evidence, gathered via microsatellite genotyping of pregnant females and their embryos, suggests that MP is very frequent in this species. Knowledge of MP in P. glauca should help describe more precisely its reproductive biology and contribute to the management of its populations.     

Genetic polyandry and sexual conflict in the sandbar shark, Carcharhinus plumbeus, in the western North Atlantic and Gulf of Mexico

To investigate patterns of polyandry in the sandbar shark (Carcharhinus plumbeus), 20 pregnant females were sampled from the western North Atlantic and Gulf of Mexico. Five species-specific microsatellite markers were used to genotype each shark and its litter. Of 20 litters, 17 (85%) were shown to have multiple sires. In multiply sired litters, the estimated minimum number of sires ranged from two to five with an average of 2.3 males per litter. Regression analysis did not demonstrate a significant relationship between female reproductive success and female body size or sire number and female body size. There was a high incidence of reproductive skew noted in litters, and two groups of males with significantly different mean reproductive success were observed. Analyses using Bateman's principles suggest that there is less direct benefit for females that acquire multiple mates than for males who bias paternity within litters. In light of past morphological and behavioural studies, these data suggest that patterns of polyandry in elasmobranchs may be determined by coercive mating, and that breeding behaviour has likely evolved in the context of sexual conflict.     

Reproductive biology of Carcharhinus acronotus in the coastal waters of South Carolina

The reproductive biology of blacknose sharks Carcharhinus acronotus in the western North Atlantic Ocean was studied by examining specimens collected in the coastal waters of South Carolina. Males begin the maturation process between 875 and 910 mm fork length ( L _F), as indicated by the presence of functional claspers and siphon sacs. The presence of vitellogenic oocytes and developing oviducal glands and uteri indicated that females begin to mature at c . 870 mm L _F. Length at which 50% of the population reached maturity was 896 and 964 mm L _F, equivalent to 4·3 and 4·5 years, for males and females, respectively. Gonado‐somatic indices suggested that spermatogenesis and vitellogenesis began after December. Mating took place during the end of May and the beginning of June. Fertilization occurred during late June and early July, suggesting that female blacknose sharks were capable of sperm storage. Based on the timing of fertilization and occurrence of females carrying near‐term pups in late May and early June, the gestation period for blacknose sharks was c . 11 months. Female blacknose sharks reproduced biennially based on the absence of vitellogenic oocytes in near‐term females and there being no indication of vitellogenesis in postpartum females. Male blacknose sharks were capable of reproducing annually as indicated by turgid genital ducts, which were observed in all mature males collected during late May and early June.     

Age and growth of the blacknose shark, <Emphasis Type="Italic">Carcharhinus acronotus</Emphasis>, in the western North Atlantic Ocean with comments on regional variation in growth rates

We examined the age and growth of the blacknose shark, Carcharhinus acronotus, in the western North Atlantic Ocean by obtaining direct age estimates using vertebral centra. We verified annual deposition of growth increments with marginal increment analysis and validated it by analyzing vertebrae marked with oxytetracycline from a female blacknose shark held in captivity. Von Bertalanffy growth parameters indicated that female blacknose sharks have a lower growth constant (k), a larger theortical maximum size (L), and are longer lived than males. We compared these growth parameters for blacknose sharks in the western North Atlantic Ocean to growth parameters for blacknose sharks collected in the eastern Gulf of Mexico to test for differences between regions. Females in the western North Atlantic Ocean have a significantly lower L, lower k, and a higher theoretical longevity than females in the Gulf of Mexico. Males in the western North Atlantic Ocean have a higher L<>, lower k, and higher theoretical longevity than males in the Gulf of Mexico. The significant differences between these life history parameters for blacknose sharks suggest that, when possible, future management initiatives concerning blacknose sharks should consider managing the populations in the western North Atlantic and the Gulf of Mexico as separate stocks.     

Population structure,connectivity, and demographic history of an apex marine predator,the bull shark Carcharhinus leucas

Knowledge of population structure, connectivity, and effective population size remains limited for many marine apex predators, including the bull shark Carcharhinus leucas. This large‐bodied coastal shark is distributed worldwide in warm temperate and tropical waters, and uses estuaries and rivers as nurseries. As an apex predator, the bull shark likely plays a vital ecological role within marine food webs, but is at risk due to inshore habitat degradation and various fishing pressures. We investigated the bull shark's global population structure and demographic history by analyzing the genetic diversity of 370 individuals from 11 different locations using 25 microsatellite loci and three mitochondrial genes (CR, nd4, and cytb). Both types of markers revealed clustering between sharks from the Western Atlantic and those from the Western Pacific and the Western Indian Ocean, with no contemporary gene flow. Microsatellite data suggested low differentiation between the Western Indian Ocean and the Western Pacific, but substantial differentiation was found using mitochondrial DNA. Integrating information from both types of markers and using Bayesian computation with a random forest procedure (ABC‐RF), this discordance was found to be due to a complete lack of contemporary gene flow. High genetic connectivity was found both within the Western Indian Ocean and within the Western Pacific. In conclusion, these results suggest important structuring of bull shark populations globally with important gene flow occurring along coastlines, highlighting the need for management and conservation plans on regional scales rather than oceanic basin scale.     

Phylogeography and conservation of the bull shark (Carcharhinus leucas) inferred from mitochondrial and microsatellite DNA

The bull shark (Carcharhinus leucas) is a widely distributed, large coastal shark species known to travel long distances. These characteristics, coupled with the species?? long life span and late age of maturity, would lead one to predict significant global genetic exchange among bull shark populations. By contrast, data show localized depletion in some areas of large coastal shark fisheries, indicating some geographic isolation may exist. We examined genetic variation in the control region of mitochondrial DNA and at five nuclear microsatellite loci in bull sharks sampled from the western Atlantic to investigate the degree of population subdivision. The average per sample haplotype and nucleotide diversity in the mtDNA (0.51 ± 0.26 and 0.12% ± 0.12, respectively) and expected heterozygosity (0.84) in the microsatellite loci contrast sharply in having lower and higher values (respectively) relative to many other shark species. Significant structure exists between the Brazilian and all northern populations at the mtDNA control region (pairwise ??_ST > 0.8, P < 0.001), but not at the nuclear microsatellite loci. Adjacent northern populations show weak to no genetic differentiation for both markers. These results are congruent with restricted maternal gene flow between populations caused by female site fidelity to nursery areas. We estimate the current effective population size to be around 160,000 and 221,000 individuals for the southern and northern Atlantic populations, respectively. The philopatric habits and the relatively low levels of mtDNA genetic diversity observed in bull sharks must be considered in the conservation of this species. Our results indicate that effective bull shark management strategies will require local, regional, and international attention and cooperation.     

Reproductive biology of the brown smoothhound shark Mustelus henlei,in the northern Gulf of California,México

Female brown smoothhound sharks Mustelus henlei were found to reproduce annually. A mature female carried both developing oocytes in the ovary and developing embryos in the uteri concurrently for c. 1 year. A great variability in the size of embryos was recorded each month, and the maximum embryo sizes were found from late January to mid‐March. The largest oocytes in mature females were observed in mid‐March. Gestation lasted c. 10 months. A linear relationship between maternal total length (L_T) and the number of pups per litter (litter size one to 21) was estimated. Birth L_T was reached in c. 280 mm. Females and males matured at 570–660 and 550–560 mm L_T, respectively. Difference in the litter size among Californian coast (one to 10) and northern Gulf of California (one to 21) populations existed for this smoothhound shark.     

Reproductive biology of the Caribbean brooding octocoral Antillogorgia hystrix

The reproductive biology of the Caribbean gorgonian Antillogorgia hystrix was studied in the shallow‐water reefs of Cross Harbour, Great Abaco (The Bahamas) from 2009 to 2010. Antillogorgia hystrix is an internal brooder that reproduces annually. The population at Cross Harbour was gonochoric and the sex ratio was skewed toward females (~3:1). Oogenesis precedes spermatogenesis by several months, and lasts at least 9 months, with oocytes >100 μm in diameter first becoming visible in dissections of samples from February; mature oocytes are present in late October–November. The size of mature oocytes (400–900 μm in diameter) was greater than that of the spermaries, which were rarely larger than 400 μm. Brooded planulae were observed in polyps from early November to mid‐December, and planula release was observed in aquaria in December 2009, which suggests that planulation occurs continuously over this period. Planulae of A. hystrix contained dinoflagellate symbionts, presumably acquired during embryogenesis and/or by mature planulae while they were in the gastrovascular cavity of the polyp. Brooding is an uncommon reproductive strategy among Caribbean gorgonians and this is the first report of internal brooding in the genus Antillogorgia. The genus contains a number of sympatric species with different modes of reproduction, and knowledge of their reproductive biology is critical to understand their ecology and evolution.     

The biology of the finetooth shark,Carcharhinus isodon

Synopsis The finetooth shark inhabits shallow coastal waters of the western Atlantic from North Carolina to Brazil. It is common off the southeastern United States, where it spends the summer off Georgia and the Carolinas and winters off Florida. The species appears in the nursery and mating areas of South Carolina when the surface water temperature rises above 20° C in late April and early May. Both adults and juveniles are common in the shallow coastal waters of South Carolina through the summer, where they feed primarily on menhaden. The finetooth shark leaves the Carolinas in early fall and migrates southward as the surface water temperature decreases below 20° C. Females reach maturity at about 1350 mm TL. Males mature at about 1300 mm TL. The finetooth shark has consecutive, year-long ovarian and gestation cycles, like most carcharhinid sharks. Mating occurs from early May to early June. Freshly mated females bear a large spermozeugma at the base of each uterus. The spermozeugmata are large almond shaped masses of individual spermatozoa embedded in a supporting matrix. Embryos are lecithotrophic during their first fifteen weeks of development. Subsequently, the embryos establish a placental connection to the mother. Implantation occurs when the embryos measure about 130 mm or at about the fifteenth week of gestation. Gravid females carrying young 480–550 mm TL enter the shallow water nurseries off South Carolina in late May. Parturition occurs from late May to mid-June, after a gestation period of about twelve months, plus or minus two weeks. The young measure 480–580 mm TL at birth. Oocytes grow little during the gestation cycle. After parturition, a cohort of oocytes begins to develop, that will be ovulated the following May. Thus, the ovarian cycle lasts about a year, although most of the oocyte growth occurs in the months just prior to ovulation.     

Age and growth of the Australian sharpnose shark,Rhizoprionodon taylori,from north Queensland,Australia

Synopsis Age and growth were studied inRhizoprionodon taylori using specimens caught in Cleveland Bay, North Queensland, Australia. Von Bertalanffy growth parameters were estimated using three different techniques: vertebral ageing, back calculation and length frequency. Vertebrae from 138 specimens were sectioned and narrow circuli counted to estimate age. Marginal increment analysis verified that circuli were produced annually in late summer, probably as a result of stress during the mating season. The oldest female was 7 and male 6 years old. Von Bertalanffy growth parameters estimated from vertebral ageing data for males were t_O = 0.410 yr, K = 1.337, L = 652.2 mm, and for females t_O = 0.455 yr, K = 1.013 and L = 732.5 mm. Growth parameters determined by length frequency and back calculation techniques concurred with those from vertebral ageing. Growth of the 0+ age class was very rapid, averaging 140% of the size at birth in the first year. Males and females matured after only one year, the lowest age at maturity reported in the family Carcharhinidae. Annual growth increments decreased rapidly after maturity, and little growth occurred after three years.     

Reproductive biology of the Alaska skate Bathyraja parmifera,with comments on an intersexual individual

A total of 1357 specimens of Alaska skate Bathyraja parmifera were collected in the eastern Bering Sea by fisheries observers and during scientific groundfish surveys from 2003 to 2005. Male and female gonads were examined for maturity stage and seasonal reproductive timing. Based on seasonal reproductive data, including the occurrence of egg cases, ovum size, ovum number, shell‐gland width and gonado‐somatic index, this species appears to reproduce continually throughout the year. Potential effects of maternal size upon the size and number of mature oocytes were also investigated, with total length having a significant, although weak, influence on both. Morphology of a single intersexual individual encountered during the collection period is also described.     

Age,growth and reproductive biology of the silky shark,Carcharhinus falciformis,and the scalloped hammerhead,Sphyrna lewini,from the northwestern Gulf of Mexico

Synopsis The silky shark, Carcharhinus falciformis, and scalloped hammerhead, Sphyrna lewini, represent >80% of the shark by-catch of the winter swordfish/tuna longline fishery of the northwestern Gulf of Mexico. This catch represents a potential supplemental fishery, yet little is known of the life histories of the two species. This report relates reproductive biology data to age and growth estimates for 135 C. falciformis and 78 S. lewini. Unlike other regional populations, C. falciformis in the Gulf of Mexico may have a seasonal 12 month gestation period. Males mature at 210–220 cm TL (6–7 yr); females at >225 cm TL (7–9 yr). Application of age at length data for combined sexes produced von Bertalanffy growth model parameter estimates of L_∞ = 291 cm TL, K = 0.153, t₀ = −2.2 yr. Adult male S. lewini outnumbered adult females in catches because of differences in the distributions of the sexually segregated population. Males mature at 180 cm TL (10 yr); females at 250 cm TL (15 yr). von Bertalanffy parameter estimates for combined sexes of this species were L_∞ = 329 cm TL, K = 0.073, t_o = −2.2 yr.